Homo

Holy crap 2015 was a big year for fossils. And how fortuitous that 2016 begins on a Fossil Friday – let’s recap some of last year’s major discoveries.

Homo naledi

Some Homo naledi mandibles in order from least to most worn teeth.

The Homo naledi sample is a paleoanthropologist’s dream – a new member of the genus Homo with a unique combination of traits, countless remains belonging to at leasta dozen individuals from infant to old adult, representation of pretty much the entire skeleton, and a remarkable geological context indicative of intentional disposal of the dead (but certainly not homicide, grumble grumble grumble…). The end of 2015 saw the announcement and uproar (often quite sexist) over this amazing sample. You can expect to see more, positive things about this amazing animal in 2016.

We’ll be presenting a bunch about Homo naledi at this year’s AAPA meeting in Hotlanta. I for one will be discussing dental development at Dinaledi- here’s a teaser:

As long as we’re talking about the AAPA meetings, my colleague David Pappano and I are organizing a workshop, “Using the R Programming Language for Biological Anthropology.” Details to come!

Lemur graveyard

Homo naledi wasn’t the only miraculously copious primate sample announced in 2015. Early last year scientists also reported the discovery of an “Enormous underwater fossil graveyard,” containing fairly complete remains of probably hundreds of extinct lemurs and other animals. As with Homo naledi, such a large sample will reveal lots of critical information about the biology of these extinct species.

Australopithecus deyiremeda

We also got a new species of australopithecus last year. Australopithecus deyiremeda had fat mandibles, a relatively short face (possibly…), and smaller teeth than in contemporaneous A. afarensis. One tantalizing thing about this discovery is that we may finally be able to put a face to the mysterious foot from Burtele, since these fossils come from nearby sites of about the same geological age. Also intriguing is the possible evidence, based on published CT images (above), that A. deyiremeda had relatively advanced canine and delayed molar development, a pattern generally attributed to Homo and not other australopithecines (if this turns out to be the case, you heard it here first!).

Lomekwian stone tool industry

Roughly contemporaneous with A. deyiremeda, Harmand et al. (2015) report the earliest known stone tools from the 3.3 million year old site of Lomekwi 3 in Kenya. These tools are a bit cruder and much older than the erstwhile oldest tools, the Oldowan from 2.6 million years ago. These Lomekwian tools, and possible evidence for animal butchery at the 3.4 million year old Dikika site in Ethiopia (McPherron et al. 2010; Thompson et al. 2015), point to an earlier origin of lithic technology. Fossils attributed to Kenyanthropus platyops are also found at other sites at Lomekwi. With hints at hominin diversity but no direct associations between fossils and tools at this time, a lingering question is who exactly was making and using the first stone tools.

Earliest Homo

The reconstructed Ledi Geraru mandible (top left), compared with Homo naledi (top right), A. deyiremeda (bottom left), and the Uraha early Homo mandible from Malawi (bottom right). Jaws are scaled to roughly the same length from the front to back teeth; the Uraha mandible does not have an erupted third molar whereas the others do and are fully adult.

Just as Sonia Harmand and colleagues pushed back the origins of technology, Brian Villmoare et al. pushed back the origins of the genus Homo, with a 2.7 million year old mandible from Ledi Geraru in Ethiopia. This fossil is only a few hundred thousand years younger than Australopithecus afarensis fossils from the nearby site of Hadar. But the overall anatomy of the Ledi Geraru jaw is quite distinct from A. afarensis, and is much more similar to later Homo fossils (see image above). Hopefully 2016 will reveal other parts of the skeleton of whatever species this jaw belongs to, which will be critical in helping explain how and why our ancestors diverged from the australopithecines. (note that we don’t yet have a date for Homo naledi – maybe these will turn out to be older?)

The earlier hominin fossil record wasn’t the only part to be shaken up. A small molar (KNM-ER 51261) and a set of associated hip bones (KNM-ER 5881) extended the lower range of size variation in Middle and Early (respectively) Pleistocene Homo. It remains to be seen whether this is due to intraspecific variation, for example sex differences, or taxonomic diversity; my money would be on the former.

At the later end of the fossil human spectrum, researchers also announced an archaic looking mandible dredged up from the Taiwan Straits, and a more modern-looking brain case from Israel. The Penghu 1 mandible is likely under 200,000 years old, and suggests a late survival of archaic-looking humans in East Asia. Maybe this is a fossil Denisovan, who knows? What other human fossils are waiting to be discovered from murky depths?

The Manot 1 calvaria looks very similar to Upper Paleolithic European remains, but is about 20,000 years older. At the ESHE meetings, Israel Hershkovitz actually said the brain case compares well with the Shanidar Neandertals. So wait, is it modern or archaic? As is usually the case, with more fossils come more questions.

Crazy dinosaurs

Yi qi was bringing Skeksi back, and its upper limb had a wing-like shape not seen in any other dinosaur, bird or pterosaur. There were a number of other interesting non-human fossil announcements in 2015 (see here and here), proving yet again that evolution is far more creative than your favorite monster movie makers.

What a year – new species, new tool industries, new ranges of variation! 2015 was a great year to be a paleoanthropologist, and I’ll bet 2016 has just as much excitement in store.

We’re learning about the divergence between robust Australopithecus and early Homo 2.5-ish million years ago in my Human Evolution class this week. Because of this multiplicity of contemporaneous species, when scientists find new hominin fossils in Early Pleistocene sites, a preliminary question is, “What species is it?”

To help my students learn how we know whether certain fossils belong to the same species, and to which group new fossils might belong, in this week’s lab we compared tooth sizes of Australopithecus boisei and early Homo. After seeing how tooth sizes differed between these groups, students then tested whether they could determine whether two “mystery” fossils (KNM-ER 60000 and 62000; Leakey et al. 2012) belonged either group.

Early Pleistocene hominin fossils from Kenya. Left to right: KNM-ER 406, ER 62000 and ER 1470. At the center is one f the lab’s “mystery jaws.”

The first purpose of this lab was to help familiarize students with skull and tooth anatomy of early Pleistocene humans. Although lectures and readings are full of images, a lab activity forces students to spend time visually examining fossils. Plus, they’re in 3D which is a whole D greater than 2D – the visual equivalent of going to eleven! The second goal of the lab was to help prepare students for their term projects, in which they must pose a research question about human evolution, generate predictions, and find and use data to test hypotheses.

If you’re interested in using or adapting this activity for your class, here are the handout and data sheet into which students enter their measurements. The data sheet specifies the fossils that can be downloaded from africanfossils.org. Some relevant fossils (i.e., KNM WT 15000 and ER 992) were not included because the 3D scans yield larger measurements than in reality.

Figure 3 from Ward et al. 2015. A little distal to the hip, yes, but the pun still works. Views are, going clockwise starting at the top the top left, from above, from below, from the back, from the side, and from the front.

There’s also a partial ilium associated with the femur – that makes a pretty complete hip!

Figure 5 from Ward et al. shows the fossil. Jump for joy that it’s complete enough for us to tell it comes from the left side!

Despite how fragmentary the femur and ilium are, the researchers were able to estimate the diameter of the femur head and hip socket reliably. The hip joints are smaller than all Early Pleistocene Homo except for the Gona pelvis. Comparing ER 5881 the large contemporaneous KNM-ER 3228 hip bone, the authors found these two specimens to be more different in size than is usually seen between sexes of many primate species. The size difference best matches male-female differences in highly dimorphic species like gorillas.

Ward et al. find that the specimen generally looks like early Homo but that the inferred shape of the pelvic inlet is a little different from all other Early and Middle Pleistocene human fossils. The authors take this discrepancy to suggest that there was more than one “morphotype” (‘kind of shape’), and therefore possibly species, of Homo around 1.9 million years ago. While I wouldn’t just yet go so far as to say this anatomy is due to species differences, I do agree that KNM ER 5881 helps our understanding and appreciation of anatomical variation in our early ancestors. Like all great fossil discoveries, the more we find, the more we learn that we don’t know. Here’s to more Homo hips in the near future!

This was a big week for Middle-Late Pleistocene fossil humans. Chun-Hsiang Chang and colleagues describe a mandible dredged up off the western coast of Taiwan, which they note in the title as, “The first archaic Homo” fossil known from the region. The geological context makes it difficult to date the specimen precisely, but authors argue it is probably younger than 190 thousand years old.

The Penghu mandible. Figure 3. From Chang et al.

In life, this individual was fully grown but appears never to have developed third molars (the “wisdom teeth”). Such “third molar agenesis” is relatively rare before modern times, but is also seen in the D2735Homo erectus mandible from Dmanisi. I wouldn’t make much of this coincidence, but it does raise the question of whether the cause of agenesis, not uncommon today, was the same then as now.

The Manot calvaria (Figure 2 from Hershkovitz et al.) The views are (a-d) from the top with front to the left; from the left; from the front; and from the back. Extra credit: In the top view (a), can you identify the features telling that the front is to the left?

John Hawks has good posts dedicated to both Penghu and Manot. The upshot of these discoveries is that Middle and Late Pleistocene human population diversity, and the interactions between these populations, are probably much more complicated and interesting than the old model of ‘modern’ humans arising singly in Africa and replacing ‘archaic’ humans in different parts of the globe. With the technological advances and fossil discoveries of the past decade, the rather simple Replacement model has given way to a better appreciation of true complexity of human evolution toward the end of the Ice Age. Both of these new papers reflect this new perspective.

Along these lines, accompanying the Manot paper in Nature is an editorial, “Human history defies easy stories.” What caught my attention reading this (anonymous?) commentary is that it puts scientific interpretations of the past into a social and historical context. The author notes that the traditional story of modern humans arising, spreading and eradicating other groups of human has “imperialist framing, in which evolution and replacement can be justified after the fact as a kind of manifest destiny.” Science doesn’t occur in a vacuum, it’s done by people whose minds and creativities are molded in specific historical, economic and cultural contexts. This editorial comment makes one wonder how the human fossil record would have been interpreted, had most of it not discovered against the social backdrop of ruthless capitalism.

It’s been a quiet month here at Lawnchair, as I’ve just returned from the Rising Star Workshop, taking part in the analysis and description of new hominin remains from South Africa. We’ll have some exciting announcements to make in the near future.

Also, I petted a ferocious, bloodthirsty lion!

To ease back into the Lawnchair, I thought I’d resurrect eFfing Fossil Friday, a short-lived series from when I was collecting data for my dissertation three years ago (speaking of which, a paper related to my dissertation came out in AJPA during the Workshop, as well). A lot has happened since the last installment of FFF, so whose heads will be on the chopping block today?

Crania #s 9, 15 and 17 (clockwise from top left). Cranium 9 is an early adolescent and the other two are adults – lookit how the facial anatomy changes with age! (Fig. 1 from Arsuaga et al., 2014)

It’s new crania from Sima de los Huesos, Atapuerca! These are published today in the journal Science by Juan L. Arsuaga and colleagues. Sima de los Huesos is a pretty remarkable site in Spain dating to the Middle Pleistocene; the site is probably at least 400,000 years old, and the remains of at least 28 individuals. These specimens show many similarities with Neandertals who later inhabited the area, but don’t have all of the ‘classic’ Neandertal features.

What I like about this figure from the paper is that the comparison of the adolescent (top left) with adults (the other two) shows how the skull changes during growth. The major visible difference is that the face sticks out in front of the brain case more in the adults than the adolescent. As a result, the adolescent lacks a supraorbital torus (“brow ridge”), but this would have developed as the face grew forward and away from the brain. Ontogeny!

A few years ago, Scott Simpson and colleagues published some of the most complete fossil human hips (right). The fossils are from the Busidima geological formation in the Gona region of Ethiopia, dated to between 0.9-1.4 million years ago. (Back when I wasn’t the only author of this blog, my friend and colleague Caroline VanSickle wrote about it here)

Researchers attributed the pelvis to Homo erectus on the basis of its late geological age and a number of derived (Homo-like) features. In addition, the pelvis’s very small size indicated it probably belonged to a female. One implication of this fossil was that male and female H. erectus differed drastically in body size.

Christopher Ruff (2010) took issue with how small this specimen was, noting that its overall size is more similar to the small-bodied Australopithecus species. Using the size of the hip joint as a proxy for body mass, Ruff argued Gona’s small size would imply a profound amount of sexual dimorphism in H. erectus: much higher than if Gona is excluded from this species, and higher than in modern humans or other fossil humans. Ruff thus proposed an alternative hypothesis to marked sexual dimorphism, that the Gona pelvis may have belonged to an australopithecine.

Fig. 3 From Ruff’s (2010) reply. Australopiths (and Orrorin) are squares and Homo are circles. Gona’s estimated femur head diameter is represented by the star and bar.

Now, Simpson & team replied to Ruff’s comments, providing a laundry list of reasons why this pelvis is H. erectus and not Australopithecus. They cite many anatomical features of the pelvis shared with Gona and Homo fossils, but not australopithecines. They also note that there are many other bones reflective of body size, that seem to suggest a substantial amount of size variation in Homo fossils, even those from a single site such as Dmanisi (Lordkipanadze et al., 2007).

Interestingly, neither of these parties compared the implied size variation with that of living apes. So I’ll do it! Now, I do not have any acetabulum data, but a friend lent me some femur head measurements for living great apes a few years ago. Gona is a pelvis and not a femur, but there are more fossil femora than hips. Because there’s a very high correlation between femur head and acetabulum size, Ruff estimated Gona’s femur head diameter to be 32.6 mm (95% confidence interval: 30.1-35.2; Simpson et al. initially estimated 35.1 mm based on a different dataset and method). To quantify size variation, we can compare ratios of larger femur heads divided by smaller ones. Now, this ratio quantifies inter-individual variation, but it will underestimate sexual dimorphism since I’m likely sampling some same-sex pairs that aren’t so different in size. But this is just a quick and dirty look. So, here’s a box plot of these ratios for Homo fossils, larger specimens divided by Gona’s estimated femur head size in different time periods:

Clearly, Gona is much smaller than most other fossil Homo hips, since ratios are never smaller than 1.14. Average body size increases over time in the Homo lineage, reflected in increasing ratios from left to right on the plot. Early Pleistocene Homo fossils are fairly small, including Dmanisi, hence the lower ratios than later time periods. Middle Pleistocene Homo (MP), represented by the most fossils, shows a large range of variation, but even the smallest is still 1.17 times larger than the largest estimate of Gona’s femur head size. To put this into context, here are those green ratios (assuming a larger size for Gona) compared with large/small ratios from resampled pairs of living apes and humans:

The fossil ratios of larger/smaller HD from above, compared with resampled ratios from unsexed living apes and humans. Boxes include the 50% quartiles, and the thick lines within are sample medians. **(05/03/14: This plot has been modified from the original version post, which only included the fossil ratios based on the smaller Gona estimate)

What we see for the extant apes and humans makes sense: humans and chimpanzees show smaller differences on average, whereas average differences between gorillas and orangutans are larger. This accords with patterns of sexual dimorphism in these species. **What this larger box plot shows is that if we accept Ruff’s smaller average estimate of Gona’s femur head size (white boxes), it is relatively rare to sample two living specimens so different in size as seen between Gona and other fossils. If we use Simpson et al.’s larger Gona size estimate, variation is still elevated above most living ape ratios. Only when Gona is compared with the generally-smaller, earlier Pleistocene fossils, does the estimated range of variation show decent overlap with living species. Even then, the overlap is still above the median values.

These results based on living species agree with Ruff’s concern, that including Gona in Homo erectus results in an unusually large range of variation in this species. Such a large size range isn’t necessarily impossible, but it would be surprising to see more variation than is common in gorillas and orangutans, where sexual size dimorphism is tremendous. Ruff suggested that the australopith-sized Gona pelvis may in fact be an australopith. This was initially deemed unlikely, in part because the fossil is well-dated to relatively late, 0.9-1.4 million years ago. However, Dominguez-Rodgrigo and colleauges (2013) recently reported a 1.34 mya Australopithecus boisei skeleton from Olduvai Gorge, so it is possible that australopiths persisted longer than we’ve got fossil evidence for, and Gona is one of the latest holdouts.

So many possible explanations. More clarity may come with further study of the fossils at hand, but chances are we won’t be able to eliminate any of these possibilities until we get more fossils. (also, the post title wasn’t a jab at the fossils or researchers, but rather a reference to the movie Office Space)

It’s a new year, and while my mind should be marred by a hangover, instead all I can think about are fossils and scapulas.

A pretty cool study was published online in the Journal of Human Evolution last week, and I’ve finally gotten to peruse it. Fabio Di Vincenzo and colleagues analyzed the shape of the outline of the glenoid fossa on the scapula (not to be confused with the glenoid on your skull), from Australopithecus africanus to present day humans. The glenoid fossa is essentially the socket in the ball-and-socket joint of your shoulder. The authors found that there is pretty much a single trend of glenoid shape change from Australopithecus through the evolution of the genus Homo: from the fairly narrow joint in Australopithecusafricanus and A. sediba, to the relatively wide joint in recent humans. The overall size and shape of the joint influences/reflects shoulder mobility, so presumably this shape change hints that more front-to-back arm motions became more important through the course of human evolution (authors suggest throwing in humans from the Late Pleistocene onward).

The finding of a single predominant trend in glenoid shape evolution is pretty interesting. On top of that, the authors add an ‘evo-devo’ twist by comparing species’ average “shapes” (first principle component scores, on the y-axis in the figure at right) with their estimated ages at skeletal maturity (which appears scaled to the modern human value, on the x-axis). Though it’s not an ideal dataset for running a linear regression, the figure at right shows that there appears to be a fairly linear relationship across human evolution, such that groups with an older age at skeletal maturity tend to have a more rounded (modern human-like) glenoid fossa (note that the individuals in the analysis were all adults). Overall size does not contribute to shape variation among these glenoids.

This work raises two issues, and ultimately leads to a testable evo-devo hypothesis. The first issue is to what extent we can trust their estimates of age at skeletal maturity. These estimates were allegedly taken from a chapter by Helmut Hemmer (2007) in the prohibitively expensive Handbook of Paleoanthropology. Cursorily glancing at this chapter, I can’t find age at skeletal maturation estimated for any hominids. It is possible that in my skimming I missed the estimates, or alternatively that Di Vincenzo and colleagues misinterpreted another variable as skeletal development. Either way, these estimates would still need to be taken with a grain of salt, given that it is almost impossible to know the true age at death of a fossil (but see Antoine et al. 2008), especially if there are no associated cranio-dental elements.

That said, it is perfectly reasonable to suppose that the age at skeletal maturation has increased over the course of human evolution; life-span increased through human evolution, and so all else being equal (which it almost certainly isn’t) we could expect that maturation would occur later over time, too. So this leads to a second issue: given the “evo-devo change” the authors hypothesize, what is the evo-devo mechanism? That is, how was development modified to effect the evolutionary changes we see in the hominid scapula? Because they found adult glenoid shape correlates with estimated age at skeletal maturity, this leads to the hypothesis that postnatal skeletal growth accounts for the shape difference. Indeed, they state:

“If functional and static allometric influences are unlikely, we…interpret the trend…as reflecting growth and developmental factors. A major, albeit gradual, trend of ontogenetic heterochrony occurred in the evolution of the genus Homo… and thus differences within and between taxa in overall growth rates may have produced the pattern of variation between samples, as well as the overall temporal trend observed. The regression of life history variables [they only looked at 1]… with PCA [principle components analysis] scores supports this ‘ontogenetic’ hypothesis.”

The authors suggest that humans’ slower growth rates but longer growth period “led to longer periods of bone deposition along the inferior-lateral edge of the [glenoid fossa]” The heterochronic process they suggest is “peramorphosis” – the descendant reaches a shape that is ‘beyond’ that of the ancestor.

The figure above is from a seminal “heterochrony” paper by Pere Alberch and colleagues (1979), portraying how peramorphosis can occur. In each, the y-axis represents shape and the x-axis is age. A the descendant’s peramorphic shape (“Ya”) could result from accelerated growth (left graph) or from an extension of growth to later ages than in the ancestor (right graph).

And so this leads to a testable hypothesis. Di Vincenzo and colleagues cite (dental) evidence that humans’ overall body growth rates are slower than earlier hominids’, undermining the hypothesis that acceleration is responsible for humans’ glenoid peramorphosis. Rather, they hypothesize that humans’ slower growth rates coupled with a longer period of skeletal development, to result in a relatively wider glenoid, due to increased development of the secondary growth centers (e.g. the graph at right, above). This developmental scenario predicts that subadult human glenoids should resemble earlier hominid adults’, that “ontogeny recapitulates phylogeny” as far as glenoid shape is concerned. Analyzing glenoid growth can even be extended to include fossils – the >3 million year old human ancestor Australopithecus afarensis has glenoids preserved for an infant (DIK-VP-1; Alemseged et al. 2006) and 2 adults (AL 288 “Lucy” and KSD-VP-1; Johanson et al. 1982, Haile-Selassie et al. 2010). An alternate hypothesis is that species’ distinct glenoid shapes are established early during life (i.e. in utero), and/or that no simple heterochronic process is involved.

Di Vincenzo’s and colleagues’ study points to the importance of development in understanding human evolution, and their hypothesized “evo-devo change” in glenoid shape is ripe for testing.

Antoine, D., Hillson, S., & Dean, M. (2009). The developmental clock of dental enamel: a test for the periodicity of prism cross-striations in modern humans and an evaluation of the most likely sources of error in histological studies of this kind Journal of Anatomy, 214 (1), 45-55 DOI: 10.1111/j.1469-7580.2008.01010.x